Scientia Horticulturae, 19 (1983) 311--319 311

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

HISTOLOGICAL AND ANATOMICAL CHARACTERISTICS OF

PARTHENOCARPIC AND NORMAL PEAR FRUITS

M. CLARA MARCUCCI' and T. VISSER 2

' Istituto Coltivazioni Arboree, Universitd di Bologna (Italy)
2Institute for Horticultural Plant Breeding, Wageningen (The Netherlands)
(Accepted for publication 8 July 1982)

ABSTRACT

Marcucci, M.C. and Visser, T., 1983. Histological and anatomical characteristics of par-
thenocarpic and normal pear fruits. Scientia Hortic., 19: 311--319.

The external and internal development of normal and parthenocarpic 'Bonne Louise'
and 'Conference' pear fruits was studied. The diameter of the normal fruits and their lo-
cules increased almost linearly between mid-April (bloom) and the end of June. There-
after, the fruits continued to grow, while locule expansion virtually stopped. Partheno-
carpic fruits grew at nearly the same rate as normal ones, although the length/diameter
ratio was consistently larger. The mean width of their locules had already begun to de-
crease from about mid-May because undifferentiated tissue started to grow inside, in due
course partly or entirely filling the locules.
In 'Conference', this led to the disappearance of locules and core in many fruits; in
'Bonne Louise', the locules remained visible. In addition, the pith tissue surrounding the
carpels appeared to exert pressure on the locules, with the result that they became partly
or entirely flattened in 'Bonne Louise' and 'Conference' fruits, respectively. In the normal
fruits, locules with normal seeds were much larger than locules with only abortive seeds.
The parthenocarpic fruits had a much smaller core than the normal fruits with abortive
seeds, the core of the latter being smaller than in fruits with normal seeds. Fruit diameter
decreased with core diameter; on this account parthenocarpic pear fruits were more slen-
der than normal fruits.

INTRODUCTION

T h e o c c u r r e n c e o f seedless fruits is c o m m o n in m a n y pear cultivars. Al-

t h o u g h t h e t e r m p a r t h e n o c a r p y is o f t e n e m p l o y e d w h e n seedlessness is
m e a n t ( G o r t e r a n d Visser, 1 9 5 8 ) , in t h e strict sense it signifies the develop-
m e n t o f fruits w i t h o u t seeds or w i t h seeds w i t h o u t e m b r y o s . E i t h e r t e r m ap-
plies in t h e case o f spring f r o s t d u r i n g b l o o m , w h i c h d e s t r o y s style and ova-
ries just b e f o r e or a f t e r p o l l i n a t i o n . P a r t h e n o c a r p i c or seedless fruits usually
differ in shape, having a relatively smaller d i a m e t e r t h a n fruits with seeds, as,
f o r e x a m p l e , o b s e r v e d b y S c h a n d e r ( 1 9 5 5 ) f o r ' C o n f e r e n c e ' and Griggs et al.
( 1 9 5 7 ) f o r ' B a r t l e t t ' pear. In a d d i t i o n , Staritski ( 1 9 7 0 ) o b s e r v e d t h a t the

0304-4238/83/$03.00 © 1983 Elsevier Science Publishers B.V.

312

core of seedless fruits of P y r u s nivalis was smaller than that of fruits with
seeds.
For 'Conference' and 'Beurr~ Hardy' fruits, we have frequently noted that
no core at all appears to be present. In view of the lack of information on
the latter aspect, our aim was to investigate the development of both seeded
and seedless pear fruits, with special attention to the core and locule.

MATERIAL AND METHODS

The observations were carried out in an orchard at Elst on the pear culti-
vars 'Conference' and 'Bonne Louise d'Avranches'. Fruits were collected at
various times between 1 and 80 days after the start of bloom and at harvest
time. At the end of bloom, which lasted about 10 days, a heavy night frost in
the unprotected part of the orchard severely damaged the flowers; styles
blackened, and usually the ovaries also. In another area, protected by over-
head sprinklers, damage was negligible. It was therefore possible to collect
both normal and frozen, parthenocarpic, fruits.
In an early stage of development, the fruits were cut with a dry-ice micro-
tome, and the 40-gthick slices were bleached in a 2% chlorine solution and
stained with acid fuchsine in lactophenol. Later, when the fruits were larger
and tougher (due to the presence of scleroids), they were cut transversely
with a knife.
The following measurements were made on 8--14 fruits per date and per
category:
F r u i t d i a m e t e r = the greatest cross-sectional diameter of the fruit.
F r u i t length = overall length of the fruit.
Core d i a m e t e r = the diameter of the circle drawn through the 5 dorsal
carpellary bundles at the apex of the carpels in a cross-section of fruit (Tukey
and Young, 1942). This area is a little smaller than t h a t defined by the core
line (Tukey and Young, 1942) which is drawn through the ring of sepal and
petal vascular bundles separating the p i t h from the c o r t e x (Bain, 1961). In
this context, the core is made up of the p i t h + carpels, the latter consisting
of locules and carpel walls.
L o c u l e width, measured at the widest part in cross-section.

RESULTS

For a period of 80 days after bloom, the relationship between fruit diam-
eter and time could be presented linearly (r = 0.99***, n = 14). The regres-
sions for parthenocarpic and normal fruits of 'Conference' were y = 2.30 +
0.26x and y = 1.26 + 0.31x, respectively (y = diameter, x = days after bloom)
and diverged with time, indicating that the diameter of parthenocarpic fruits
increased at a somewhat lower rate than t h a t of normal fruits. As the fruit
length of both fruit types increased similarly, the length]diameter ( L / D )
ratio of the parthenocarpic fruits tended to be larger than that of the normal
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TABLE I

M e a s u r e m e n t s in m m b a s e d o n 8 - - 1 2 pear fruits w i t h a similar average l e n g t h ( L ) p e r d a t e

a n d p e r c a t e g o r y : f r u i t d i a m e t e r (D), l e n g t h / d i a m e t e r {L/D) r a t i o , a n d locule w i d t h w h e n
seeds w e r e a b s e n t or a b o r t i v e ( A B ) o r n o r m a l ( N O R ) in small a n d large, p a r t h e n o c a r p i c
and normal fruits

Date Fruit category 'Bonne Louise' 'Conference'

and condition
Fruit LID Locule width Fruit LID Locule width
diam- ratio w h e n seeds diam- ratio w h e n seeds
eter eter
AB l NOR AB NOR

18 J u n e Small, parth. 16.2 2.02 0.72 -- 14.3 2.56 0.37 --

Small, normal 19.4 1.82 1.90 2.78 18.6 2.41 1.19 2.53
Large, parth. 27.1 1.65 0.71 -- 21.8 2.42 0.80 --
Large, no~mal 27.2 1.66 1.94 3.28 24.3 2.14 1.51 3.16

17 J u l y Small, parth. 25.7 1.85 0.94 -- 25.7 2.26 0.47 --

Small, normal 32.2 1.53 2.26 4.25 30.2 2.01 1.21 3.06
Large, parth. 35.0 1.61 0.86 -- 30.6 2.22 0.41 --
Large, normal 38.3 1.50 2.78 4.62 36.0 1.96 1.24 3.02

24 S e p t . Small, parth. 52.0 1.59 0.82 -- 50.3 2.26 2 _

Small, normal 56.2 1.44 2.56 4.07 53.5 2.00 1.40 4.30
Large, parth. 62.3 1.47 0.42 -- 62.7 2.14 3 _
Large, normal 63.4 1.48 2.40 4.50 70.0 1.96 1.72 4.28

'T h e smaller " a b o r t i v e " seeds (see Fig. 4) were still w h i t e in J u n e ; l a t e r o n t h e y t u r n e d

b r o w n inside, shrivelled a n d e n d e d u p as f l a t t e n e d i n t e g u m e n t s at h a r v e s t .
2 N e a r l y all o f t h e small fruits w e r e d e v o i d o f locules a n d n o core was discernible.
3 E i g h t y p e r c e n t o f t h e locules were f l a t t e n e d a n d t h e i r w i d t h was negligible.

26 Conference Bonne Louise

o
c

\/
"6

~
L 14

08 I~#'O
.si. •

ii /
o
/

0215/4 1/5 15/5 1/6 15/6 7 15/4 1/5 15/5 1 6 15/6 1 7

(1) (17) (31) (48) (62) (78) (5) (21) (35) (52) (16) (82)
Date (days after start of bloom )

Fig. 1. Relation between time (days after start of bloom) and the mean diameter of par-
thenocarpic pear fruits and that of their locules.
314

fruits (Table I). The ratio was also usually larger for the smaller than for the
larger fruits, which, as regards the normal fruits, corresponded with the
smaller fruits having fewer seeds. The ratio also became smaller as the fruits
became larger with time (Table I).
For the period from mid-April, when bloom started, until the end of June,
locule width of the normal fruits showed a linear increase with time (r =
0.93***, n = 11--14), similar to the fruit diameter. Between June and the
end of September (harvest), fruit diameter (and length) continued to increase
sharply, but the locules widened only slightly (Table I). The picture for the
locules of the parthenocarpic fruits was quite different. The locule diameter
increased only until about mid-May, and thereafter decreased, in contrast
with the linearly increasing fruit diameter (Fig. 1). This was owing to the
fact that some 5 weeks after bloom, undifferentiated tissue started growing
in m a n y locules, which were in due course partly or entirely filled (Figs. 2
and 3). In various cases, this tissue appeared to originate from outside the
carpel wall which was penetrated by the tissue, continuing to grow thereafter.
The tissue sometimes originated from the carpel wall itself. In the normal
fruits, tissue growing in the locules was never seen with 'Bonne Louise' and
rarely with 'Conference'.
A second factor influencing locule size appears to be the presence or ab-

Fig. 2. Undifferentiated tissue growing in the locules of parthenocarpic fruits of 'Bonne

Louise' (left) and 'Conference' (right) on 26 May, about 6 weeks after bloom started.
 315

Fig. 3. Cross s e c t i o n s on 3 0 J u n e , a b o u t 11 w e e k s after b l o o m started, o f parthenocarpic

fruits o f ' B o n n e Louise' ( l e f t ) - - s h o w i n g that the l o c u l e s m a i n t a i n e d their shape and are
more or less filled w i t h tissue - - and o f ' C o n f e r e n c e ' (right) - - s h o w i n g that the l o c u l e s are
partly f l a t t e n e d , o n e o f t h e m being filled w i t h tissue.

sence of seeds. Table I and Fig. 4 show that the width of locules in the nor-
mal fruits, containing small-sized abortive seeds only, was half or less than
half that of locules with the larger normal seeds. Generally, the locules of the
larger fruits were only slightly wider than those of the smaller fruits. Due to
the presence of the tissue inside or due to pressure from the outside, the lo-
cules of the totally seedless, parthenocarpic, fruits were very much narrower
than those with abortive seeds in the case of normal fruits. By early June, an
average of 2 of the 5 locules of parthenocarpic fruits of 'Bonne Louise' con-
tained such tissue. This increased to virtually all locules having tissue at har-
vest (24 September), but the mean locule width changed little. In partheno-
carpic 'Conference' fruits, the percentage of locules either closed by tissue or
flattened (Fig. 5, left), or n o t discernible (Fig. 5, right) increased from 30 in
mid.June, 43 at the end of June, and 54 in mid~luly to 90 at harvest. This in-
crease was partly due to an increase of fruits in which no core, and thus no
locules, could be seen: from 17% in June to 41% in July and September. In
the case of " f l a t t e n e d " locules, the tissue formed inside was also compressed.
The smaller diameter of the parthenocarpic 'Bonne Louise' fruits was ac-
companied by a smaller core as compared with the normal fruits in the same
category (Table II). Similarly, the coreless 'Conference' fruits (see Fig. 5)
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Fig. 4. Cross section on 30 June, about 11 weeks after bloom, of a normal fruit of 'Con-
ference' pear showing in each of the locules left and right a normal seed flanked by a
smaller "abortive" seed; both the latter and the small seeds in the locules above and below
are still white, but will turn brown and shrivel later on; note that locules with good seeds
are larger than those with poor ones.

TABLE II

Mean length (L), diameter (D) and ratio (L/D) of fruits, diameter of core (C) and cortex
(D--C) of different categories of 'Bonne Louise' and 'Conference' fruits at harvest; means
in m m of 7--10 fruits per category. Cortex width as a percentage of the fruit diameter
(D) in parentheses

Fruit category L D L/D Core Cortex

and condition (C) (D--C)

'Bonne Louise'
Parth., small 81.7 52.4 1.56 8.0 44.4 (87%)
Normal, small 80.7 56.7 1.42 13.6 43.1 (76%)
Parth., large 96.1 61.5 1.56 9.1 52.3 (85%)
Normal, large 96.0 64.1 1.52 15.4 48.7 (76%)

'Conference'
Parth., no core 112.2 50.1 2.24 0 50.1 (100%)
Parth., with core 116.0 56.9 2.04 9.3 47.3 (83%)
Normal, aborted seeds 114.3 56.5 2.02 14.1 42.2 (75%)
Normal, good seeds 114.8 66.0 1.73 20.2 45.8 (69%)
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Fig. 5. In July, about 14 weeks after bloom, locules of many parthenocarpic fruits of
'Conference' pear had become compressed, but the undifferentiated tissue is still visible
(left). At a further stage (right), locules and core had disappeared, leaving only a hole.

had a smaller d i a m e t e r t h a n t h o s e with a core, and the fruits w i t h seeds had a

m u c h larger core t h a n t h o s e w i t h o u t ( p a r t h e n o c a r p i c ) or w i t h a b o r t e d seeds
(normal). However, the fruits with t h e a b o r t e d seeds had an a p p r e c i a b l y larger
core d i a m e t e r t h a n t h e seedless fruits. I t was also n o t e w o r t h y (Table II) t h a t
the " c o r t e x " o f t h e n o r m a l fruits was a little t h i n n e r t h a n the c o r t e x o f the
p a r t h e n o c a r p i c fruits.

DISCUSSION AND CONCLUSIONS

O u r d a t a illustrate t h a t in p e a r orchards a f t e r a spring frost, p a r t h e n o c a r p i c

o r seedless fruits result, w h i c h are m o r e slender t h a n fruits w i t h seeds. This
applied t o b o t h ' B o n n e L o u i s e ' and ' C o n f e r e n c e ' and, as the l i t e r a t u r e indi-
cates, also t o o t h e r p e a r cultivars such as ' B a r t l e t t ' (Griggs et al., 1 9 5 7 ) and
'Williams Bon Cr~tien' (Bain, 1 9 6 1 ) . G e n e r a l l y , for e x a m p l e as o b s e r v e d b y
S c h a n d e r ( 1 9 5 5 ) f o r ' C o n f e r e n c e ' , t h e l e n g t h / d i a m e t e r ratio o f t h e fruit in-
creases as the n u m b e r o f seeds per fruit decreases. F r o m this, S c h a n d e r con-
c l u d e d t h a t t h e seed p l a y an i m p o r t a n t p a r t in shaping t h e fruits. F o r an ex-
a m p l e , h e r e f e r r e d t o t h e a s y m m e t r y in fruits -- observable in the triploid
apple 'Belle de B o s k o o p ' -- d u e t o the p r e s e n c e o f a single seed o n o n e side.
O n the o t h e r h a n d , Staritski ( 1 9 7 0 ) s t a t e d t h a t " t h e seed's role as a g r o w t h
318

centre should n o t be overestimated". This statement appears to be related to

the observation that: "the unilateral position of seeds in Pyrus nivalis may
cause local enlargement of the core. Since the layer of flesh outside the core
is uniformly thick, a rather asymmetric fruit is in fact formed".
Therefore, the seeds do, in fact, shape the fruit, albeit indirectly by shap-
ing the core. This can also be derived from the observations of both Staritski
(1970) and ourselves that the core diameter of seedless fruits is much smaller
than the core diameter of fruits with seeds. Our data also show -- in keeping
with Staritski's finding on asymmetric fruits (see above) -- that the cortex
diameter is largely independent of the presence of seeds. It may therefore be
concluded that with respect to (transverse) cortex development, the seeds
play a minor role. Expressed as a percentage of fruit diameter, the cortex
diameter varied between 76 and 87 for 'Bonne Louise' and between 69 and
83 for 'Conference' fruits with a core (Table II). These percen'tages are of a
similar order as the 82--84% found for 'Williams Bon Cr~tien' by Bain (1961).
Our data further show that core size and locule size w as influenced by the
presence of seeds -- were related. Core and locules also have in c o m m o n that
their expansion, which is initially almost linear, nearly stops in July, some 12
weeks after bloom, in contrast with the continuing growth of the fruit (Tukey
and Young, 1942; this paper).
However, locule and core development in parthenocarpic fruits were also
quite different with respect to a p h e n o m e n o n on which we were unable to
find information in the literature. It concerns the growth, starting a b o u t the
third week of May, some 5 weeks from bloom, of a presumably undifferen-
tiated (callus?) tissue in the locules along the carpel wall, resulting in a de-
crease of mean locule width. As growth continues, the locules may become
completely filled with tissue in many cases. Neverthelesss, the locules re-
mained visible in 'Bonne Louise', b u t this process appeared to lead to their
disappearance in 'Conference', thus making the core no longer discernible. A
second reason for a reduction in locule size seemed to be the pressure exerted
by the pith tissue surrounding the carpels. This occurred even in the normal
fruits, as shown by the smaller size of locules containing only "abortive"
seeds compared to those with the larger normal seeds able to provide more
counter pressure. It is n o t surprising, therefore, that in the totally seedless
fruits only a line instead of the locule was visible because of complete flattening.
This occurred only in parthenocarpic 'Conference' fruits, b u t not in those of
'Bonne Louise', of Which the locules, though smaller than in the normal
fruits, remained intact. The 'Bonne Louise' locules generally contained more
tissue than the 'Conference' ones, so it is possible that the former's locules
were not completely closed because of the counter pressure exerted by the
internal tissue.
Finally, it is worth mentioning that the locule tissue of both pears grew
actively (10-fold increase in volume in 3 weeks) on media containing 0.1--1 mg
kinetin per 1. This confirms that it concerned a u t o n o m o u s l y growing tissue
in the locules and n o t residual material.
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ACKNOWLEDGEMENTS

The first author (M.C.M.) was able to participate in this work as a result of
a fellowship provided by the Italian National Research Council. We gratefully
acknowledge the able assistance of Mr. B.A. Uijtewaal in carrying o u t part of
the work.

REFERENCES

Bain, J.M., 1961. S o m e morphological, anatomical and physiological changes in the pear
fruit (Pyrus communis var. Williams Bon Cr~tien) during development and following
harvest. Aust. J. Bot., 9: 99--123.
Gorter, C.J. and Visser, T., 1958. Parthenocarpy of pears and apples. J. Hortic. Sci., 33:
207--227.
Griggs, W.H., lwakiri, B.T. and Claypool, L.L., 1957. A comparison of growth, maturity
and quality of see.d.lessand seeded Barlett pears. Proc. A m . Soc. Hortic. Sci.,78:74--84
Schander, H., 1955. Uber die Veranderlichkeit der Fruchtgestalt bei der Birnensorte 'Con-
ference'. Mitt. Obstvers. Altes Landes 9: 271--277.
Staritski,G., 1970. The morphogenesis of the inflorescence, flower and fruit of Pyrus
nivalis Jacquin var. orientalis 'Terpo'. Meded. L a n d b o u w hogesch., Wageningen, 70 (5):
91 pp.
Tukey, H.B. and Young, J.O., 1942. Gross morphology and histology of developing fruit
of the apple. Bot. Gaz., 104: 3--25.